Caryophyllales

Caryophyllales is a diverse and heterogeneous order of flowering plants that includes the cacti, carnations, amaranths, ice plants, beets, and many carnivorous plants. Many members are succulent, having fleshy stems or leaves. The betalain pigments are unique in plants of this order and occur in all its families with the exception of Caryophyllaceae and Molluginaceae.
Description
The members of Caryophyllales include about 6% of eudicot species. This order is part of the core eudicots. Currently, the Caryophyllales contains 37 families, 749 genera, and 11,620 species The monophyly of the Caryophyllales has been supported by DNA sequences, cytochrome c sequence data and heritable characters such as anther wall development and vessel-elements with simple perforations.
Circumscription
As with all taxa, the circumscription of Caryophyllales has changed within various classification systems. All systems recognize a core of families with centrospermous ovules and seeds. More recent treatments have expanded the Caryophyllales to include many carnivorous plants.
Systematists were undecided on whether Caryophyllales should be placed within the rosid complex or sister to the asterid clade. The possible connection between sympetalous angiosperms and Caryophyllales was presaged by Bessey, Hutchinson, and others; as Lawrence relates: "The evidence is reasonably conclusive that the Primulaceae and the Caryophyllaceae have fundamentally the same type of gynecia, and as concluded by Douglas (1936)(and essentially Dickson, 1936) '...the vascular pattern and the presence of locules at the base of the ovary point to the fact that the present much reduced flower of the Primulaceae has descended from an ancestor which was characterized by a plurilocular ovary and axial placentation. This primitive flower might well be found in centrospermal stock as Wernham, Bessy, and Hutchinson have suggested.' "
Caryophyllales is separated into two suborders: Caryophyllineae and Polygonineae. These two suborders were formerly (and sometimes still are) recognized as two orders, Polygonales and Caryophyllales.
APG IV
Kewaceae, Macarthuriaceae, Microteaceae, and Petiveriaceae were added in APG IV.
APG III
As circumscribed by the APG III system (2009), this order includes the same families as the APG II system (see below) plus the new families, Limeaceae, Lophiocarpaceae, Montiaceae, Talinaceae, and Anacampserotaceae.
family Achatocarpaceae
family Aizoaceae
family Amaranthaceae
family Anacampserotaceae
family Ancistrocladaceae
family Asteropeiaceae
family Barbeuiaceae
family Basellaceae
family Cactaceae
family Caryophyllaceae
family Didiereaceae
family Dioncophyllaceae
family Droseraceae
family Drosophyllaceae
family Frankeniaceae
family Gisekiaceae
family Halophytaceae
family Kewaceae
family Limeaceae
family Lophiocarpaceae
family Macarthuriaceae
family Microteaceae
family Molluginaceae
family Montiaceae
family Nepenthaceae
family Nyctaginaceae
family Petiveriaceae
family Physenaceae
family Phytolaccaceae
family Plumbaginaceae
family Polygonaceae
family Portulacaceae
family Rhabdodendraceae
family Sarcobataceae
family Simmondsiaceae
family Stegnospermataceae
family Talinaceae
family Tamaricaceae
APG II
As circumscribed by the APG II system (2003), this order includes well-known plants like cacti, carnations, spinach, beet, rhubarb, sundews, venus fly traps, and bougainvillea. Recent molecular and biochemical evidence has resolved additional well-supported clades within the Caryophyllales.
order Caryophyllales
family Achatocarpaceae
family Aizoaceae
family Amaranthaceae
family Anacampserotaceae (added in APG III)
family Ancistrocladaceae
family Asteropeiaceae
family Barbeuiaceae
family Basellaceae
family Cactaceae
family Caryophyllaceae
family Didiereaceae
family Dioncophyllaceae
family Droseraceae
family Drosophyllaceae
family Frankeniaceae
family Gisekiaceae
family Halophytaceae
family Limeaceae (added in APG III)
family Lophiocarpaceae (added in APG III)
family Molluginaceae
family Montiaceae (added in APG III)
family Nepenthaceae
family Nyctaginaceae
family Physenaceae
family Phytolaccaceae
family Plumbaginaceae
family Polygonaceae
family Portulacaceae
family Rhabdodendraceae
family Sarcobataceae
family Simmondsiaceae
family Stegnospermataceae
family Talinaceae (added in APG III)
family Tamaricaceae
APG
This represents a slight change from the APG system, of 1998
order Caryophyllales
family Achatocarpaceae
family Aizoaceae
family Amaranthaceae
family Ancistrocladaceae
family Asteropeiaceae
family Basellaceae
family Cactaceae
family Caryophyllaceae
family Didiereaceae
family Dioncophyllaceae
family Droseraceae
family Drosophyllaceae
family Frankeniaceae
family Molluginaceae
family Nepenthaceae
family Nyctaginaceae
family Physenaceae
family Phytolaccaceae
family Plumbaginaceae
family Polygonaceae
family Portulacaceae
family Rhabdodendraceae
family Sarcobataceae
family Simmondsiaceae
family Stegnospermataceae
family Tamaricaceae
Cronquist
The Cronquist system (1981) also recognised the order, with this circumscription:
order Caryophyllales
family Achatocarpaceae
family Aizoaceae
family Amaranthaceae
family Basellaceae
family Cactaceae
family Caryophyllaceae
family Chenopodiaceae
family Didiereaceae
family Nyctaginaceae
family Phytolaccaceae
family Portulacaceae
family Molluginaceae
The difference with the order as recognized by APG lies in the first place in the concept of "order". The APG favours much larger orders and families, and the order Caryophyllales sensu APG should rather be compared to subclass Caryophyllidae sensu Cronquist.
A part of the difference lies with what families are recognized. The plants in the Stegnospermataceae and Barbeuiaceae were included in Cronquist's Phytolaccaceae. The Chenopodiaceae (still recognized by Cronquist) are included in Amaranthaceae by APG.
New to the order (sensu APG) are the Asteropeiaceae and Physenaceae, each containing a single genus, and two genera from Cronquist's order Nepenthales.
Earlier circumscriptions
Earlier systems, such as the Wettstein system, last edition in 1935, and the Engler system, updated in 1964, had a similar order under the name Centrospermae.
References
External links
Tree of Life Characteristics and Phylogenetic Relationships
Superasterids
Angiosperm orders